Relationship between Population Size and Fitness

Abstract:  Long-term effective population size, which determines rates of inbreeding, is correlated with population fitness. Fitness, in turn, influences population persistence. I synthesized data from the literature concerning the effects of population size on population fitness in natural populations of plants to determine how large populations must be to maintain levels of fitness that will provide adequate protection against environmental perturbations that can cause extinction. Integral to this comment on what has been done and what needs to be done, sThe evidence suggests that there is a linear relationship between log population size and population fitness over the range of population sizes examined. More importantly, populations will have to be maintained at sizes of >2000 individuals to maintain population fitness at levels compatible with the conservation goal of long-term persistence. This approach to estimating minimum viable population size provides estimates that are in general agreement with those from numerous other studies and strengthens the argument that conservation efforts should ultimately aim at maintaining populations of several thousand individuals to ensure long-term persistence.     

Genetic Diversity, Population Size, and Fitness in Central and Peripheral Populations of a Rare Plant Lychnis viscaria

Abstract: Genetic diversity is expected to decrease in small and isolated populations as a consequence of bottlenecks, founder effects, inbreeding, and genetic drift. The genetics and ecology of the rare perennial plant Lychnis viscaria (Caryophyllaceae) were studied in both peripheral and central populations within its distribution area. We aimed to investigate the overall level of genetic diversity, its spatial distribution, and possible differences between peripheral and central populations by examining several populations with electrophoresis. Our results showed that the level of genetic diversity varied substantially among populations (  H _exp = 0.000–0.116) and that the total level of genetic diversity (mean H _exp = 0.056) was low compared to that of other species with similar life-history attributes. The peripheral populations of L. viscaria had less genetic variation (mean H _exp = 0.034) than the central ones (0.114). Analysis of genetic structure suggested limited gene flow (mean F _ST = 0.430) and high differentiation among populations, emphasizing the role of genetic drift (  N _e m = 0.33). Isolation was even higher than expected based on the physical distance among populations. We also focused on the association between population size and genetic diversity and possible effects on fitness of these factors. Population size was positively correlated with genetic diversity. Population size and genetic diversity, however, were not associated with fitness components such as germination rate, seedling mass, or seed yield. There were no differences in the measured fitness components between peripheral and central populations. Even though small and peripheral populations had lower levels of genetic variation, they were as viable as larger populations, which emphasizes their potential value for conservation.     

Regional Consequences of Local Population Demography and Genetics in Relation to Habitat Management in Gentiana pneumonanthe

Abstract:  A joint demographic and population genetics stage-based model for a subdivided population was applied to Gentiana pneumonanthe , an early successional perennial herb, at a regional (metapopulation) scale. We used numerical simulations to determine the optimal frequency of habitat disturbance (sod cutting) and the intensity of gene flow among populations of G. pneumonanthe to manage both population viability and genetic diversity in this species. The simulations showed that even small populations that initially had near-equal allele frequencies could, if managed properly through sod cutting every 6 to 7 years, sustain their high genetic variation over the long run without gene flow. The more the allele frequencies in the small populations are skewed, however, the higher the probability that in the absence of gene flow, some alleles will be lost and within-population genetic variation will decrease even under proper management. This implies that although local population dynamics should be the major target for management, regional dynamics become important when habitat fragmentation and decreased population size lead to the loss of local genetic diversity. The recommended strategy to improve genetic composition of small populations is the introduction of seeds or seedlings of nonlocal origin.     

Genetic Variation and Outcrossing Rate in Relation to Population Size in Gentiana pneumonanthe L

The amount of genetic variation in the rare perennial herb Gentiana pneumonanthe L. was determined to explore its relation to population size. Differences in isozyme variation between maternal plants and their offspring were used to investigate the relationship between population size and outcrossing rate. In 25 populations in The Netherlands, differing in size from 1 to more than 50,000 flowering individuals, 16 allozyme loci were analyzed on leaves of maternal plants and offspring grown in a greenhouse. Population size was significantly positively correlated with the proportion of polymorphic loci, but only marginally with heterozygosity and the mean effective number of alleles. Most of the studied populations were characterized by a complete absence of rare alleles, and F -statistics suggest relatively high levels of genetic differentiation among populations and thus a low level of gene flow. Leaf samples (maternal) were mostly in Hardy-Weinberg equilibrium, while several offspring samples showed an excess of homozygotes, which suggests selection favoring heterozygotes. Because most small populations consist only of adult survivors from formerly larger populations, this may partly explain the absence of a clear relationship between genetic variation of the maternal plants and population size. A significant positive correlation was found between the level of cross-fertilization and population size. From these results, we conclude that, to some degree, small populations have a reduced level of genetic variation, while their present isolation in nature reserves has resulted in a very limited interpopulational gene flow level. At present a higher level of inbreeding in small populations contributes to a further loss of genetic variation and may also result in reduced offspring fitness.     

The Effect of Fragment Shape and Species' Sensitivity to Habitat Edges on Animal Population Size

Abstract:  Habitat fragmentation causes extinction of local animal populations by decreasing the amount of viable "core" habitat area and increasing edge effects. It is widely accepted that larger fragments make better nature reserves because core-dwelling species have a larger amount of suitable habitat. Nevertheless, fragments in real landscapes have complex, irregular shapes. We modeled the population sizes of species that have a representative range of preferences for or aversions to habitat edges at five spatial scales (within 10, 32, 100, 320, and 1000 m of an edge) in a nation-wide analysis of forest remnants in New Zealand. We hypothesized that the irregular shapes of fragments in real landscapes should generate statistically significant correlations between population density and fragment area, purely as a "geometric" effect of varying species responses to the distribution of edge habitat. Irregularly shaped fragments consistently reduced the population size of core-dwelling species by 10–100%, depending on the scale over which species responded to habitat edges. Moreover, core populations within individual fragments were spatially discontinuous, containing multiple, disjunct populations that inhabited small spatial areas and had reduced population size. The geometric effect was highly nonlinear and depended on the range of fragment sizes sampled and the scale at which species responded to habitat edges. Fragment shape played a strong role in determining population size in fragmented landscapes; thus, habitat restoration efforts may be more effective if they focus on connecting disjunct cores rather than isolated fragments.     

Genetic Diversity and Population Size in Four Rare Southern Appalachian Plant Species

Allozyme diversity was examined in four rare, high-montane plant species from the Appalachian Mountains of southeastern North America. These species may represent relictual members or descendants of an alpine community that was more widespread during the late Pleistocene. We sampled five populations of Geum radiatum (Rosaceae), Carex misera (Cyperaceae), Trichophorum cespitosum (Cyperaceae), and the four known populations of Calamagrostis cainii (Poaceae). Genetic diversity was low for all species but was typical of that found for plant species with limited ranges. Low genetic diversity may reflect historical events associated with changes in the species' biogeography. As the Pleistocene climate warmed, suitable habitat decreased in areal extent and became fragmented, probably resulting in smaller, more-isolated populations. In recent times these species, which co-occur in fragile rock outcrop habitats, have been adversely affected by human activities. Genetic analyses revealed reduced diversity in populations of decreasing size for three species. Estimates of gene flow were low ( Nm < 1.0) in all four species. Positive associations between genetic diversity and population size, evidence of recent population declines, and the low estimates of gene flow suggest that genetic drift may play a prominent role in shaping the present-day genetic composition of these species. Furthermore, these data suggest that the genetically depauperate populations are unlikely to regain genetic variation without human intervention.     

Importance of Habitat Quality and Landscape Connectivity for the Persistence of Endangered Natterjack Toads

Abstract: The natterjack toad (Bufo calamita) is endangered in several parts of its distribution, including Belgium, where it occurs mainly in artificial habitats. We parameterized a general model for natterjack population viability analysis (PVA) and tested its sensitivity to changes in the values of basic parameters. Then we assessed the relative efficiency of various conservation measures in 2 situations: a small isolated population and a system of 4 populations connected by rare dispersal movements. We based the population viability analysis on a stage‐structured model of natterjack population dynamics. We parameterized the model in the RAMAS GIS platform with vital rates obtained from our own field experience and from published studies. Simulated natterjack populations were highly sensitive to habitat quality (particularly pond drying), to dispersal from surrounding local populations, and to a lesser extent to values of fecundity and survival of terrestrial stages. Population trajectories were nearly insensitive to initial abundances, carrying capacities, and the frequency of extreme climatic conditions. The simulations showed that in habitats with highly ephemeral ponds, where premetamorphosis mortality was high, natterjack populations nearly always had a very high extinction risk. We also illustrated how low dispersal rates (<1 dispersing individual/generation) efficiently rescued declining local populations. Such source‐sink dynamics demonstrate that the identification and management of source populations should be a high priority.     

Effects of Habitat Fragmentation on Effective Population Size in the Endangered Rio Grande Silvery Minnow

Abstract:  We assessed spatial and temporal patterns of genetic diversity to evaluate effects of river fragmentation on remnant populations of the federally endangered Rio Grande silvery minnow ( Hybognathus amarus ). Analysis of microsatellite and mitochondrial DNA detected little spatial genetic structure over the current geographic range, consistent with high gene flow despite fragmentation by dams. Maximum-likelihood analysis of temporal genetic data indicated, however, that present-day effective population size ( N_eV ) of the largest extant population of this species was 78 and the ratio of effective size to adult numbers ( N_eV/N ) was ∼ 0.001 during the study period (1999 to 2001). Coalescent-based analytical methods provided an estimate of historical (river fragmentation was completed in 1975) effective size ( N_eI  ) that ranged between 10⁵ and 10⁶. We propose that disparity between contemporary and historical estimates of N_e and low contemporary N_e/N result from recent changes in demography related to river fragmentation. Rio Grande silvery minnows produce pelagic eggs and larvae subject to downstream transport through diversion dams. This life-history feature results in heavy losses of yearly reproductive effort to emigration and mortality, and extremely large variance in reproductive success among individuals and spawning localities. Interaction of pelagic early life history and river fragmentation has altered demographic and genetic dynamics of remnant populations and reduced N_e to critically low values over ecological time.     

Genetic Variation for Morphological and Allozyme Variation in Relation to Population Size in Clarkia dudleyana, an Endemic Annual

Abstract: The maintenance of genetic variation within populations is expected to allow species to respond to evolutionary challenges such as selection and environmental stress. Larger populations are generally expected to maintain larger amounts of genetic variation. Although several studies have found a positive relationship between population size and levels of genetic variation for molecular markers such as allozymes, few comparisons have been made between molecular measures of variation and genetic variation that is likely to be ecologically important. Most ecologically important traits require quantitative genetic analyses. I examined the relationship between levels of genetic variation and population size for both allozymes and morphological traits in a California endemic annual plant, Clarkia dudleyana . Levels of genetic variation for allozymes did not show a significant positive relationship with population size. The level of genetic variance for all of the 18 morphological traits exhibited no significant relationship with population size. Further, allozyme heterozygosities were not related to levels of quantitative genetic variation. These results indicate that levels of allozyme variability do not predict levels of genetic variation for morphological traits in C. dudleyana , suggesting that molecular measures of variation, in general, differ from quantitative genetic measures. These results imply that conservation genetic studies should generally focus on aspects other than measuring levels of genetic variation found within populations.     

Lineage Loss in Serengeti Cheetahs: Consequences of High Reproductive Variance and Heritability of Fitness on Effective Population Size

Abstract: In natural populations, many breeders do not leave surviving offspring, and as a result many potential genetic lineages are lost. I examined lineage extinction in Serengeti cheetahs ( Acinonyx jubatus ) and found that 76% of matrilines were lost over a 25-year period. Production of future breeders was nonrandom and generally confined to a few families. Five out of 63 matrilines accounted for 45% of the total cheetah population over the course of the study. Lineage persistence is perhaps best illustrated by the variance in lifetime reproductive success ( LRS) and heritability in this parameter. In female cheetahs, variance in LRS was high, and new data show that this LRS was heritable. Variance in LRS and heritability in LRS have dramatic consequences for effective population size, N _e. I calculated N _e for cheetahs, taking into account fluctuating population size, unequal sex ratio, non-Poisson distribution of reproductive success, and heritability of fitness. The N _e was most strongly affected by variance in reproductive success and especially heritability in reproductive success. The variance N _e was 44% of the actual population size, and the inclusion of heritability further reduced N _e to only 15% of the actual population, a ratio similar to that of a social carnivore with reproductive suppression. The current cheetah population in the Serengeti is below numbers suggested by N _e estimates as sufficient to maintain sufficient genetic diversity.     

Effects of Population Size and Food Stress on Fitness-Related Characters in the Scarce Heath, a Rare Butterfly in Western Europe

Abstract: Knowledge about the effects of inbreeding in natural populations is scarce, especially in invertebrates. We analyzed to what extent fitness-related traits in the scarce heath (  Coenonympha hero ), a butterfly, are affected by population size and isolation and whether differences in food quality influence these effects. We categorized nine populations as either large or small and isolated. Full-sib groups of offspring from 27 females were followed under seminatural conditions. Because of increased zygote mortality, egg hatchability was significantly lower in the small and isolated populations than in the large ones. Population category had no effect on larval weight under optimal conditions, but weight was significantly lower in the small-isolated category with low food quality. The effects of inbreeding can thus be hidden when conditions are benign but can appear under stress. Survival also differed significantly between population categories, and larval developmental time tended to be longer in the small-isolated category, irrespective of food conditions. We suggest that the differences in fitness between offspring from large and small isolated populations are at least partly due to inbreeding. This adds a further threat to a species that is already suffering from decreasing population sizes and increasing isolation among populations.     

Relationship of Genetic Variation to Population Size in Wildlife

Genetic diversity is one of three levels of biological diversity requiring conservation. Genetic theory predicts that levels of genetic variation should increase with effective population size. Soulé (1976) compiled the first convincing evidence that levels of genetic variation in wildlife were related to population size, but this issue remains controversial. The hypothesis that genetic variation is related to population size leads to the following predictions: (1) genetic variation within species should be related to population size; (2) genetic variation within species should be related to island size; (3) genetic variation should be related to population size within taxonomic groups; (4) widespread species should have more genetic variation than restricted species; (5) genetic variation in animals should be negatively correlated with body size; (6) genetic variation should be negatively correlated with rate of chromosome evolution; (7) genetic variation across species should be related to population size; (8) vertebrates should have less genetic variation than invertebrates or plants; (9) island populations should have less genetic variation than mainland populations; and (10) endangered species should have less genetic variation than nonendangered species. Empirical observations support all these hypotheses. There can be no doubt that genetic variation is related to population size, as Soulé proposed. Small population size reduces the evolutionary potential of wildlife species.     

Population Size and Reproduction in Phlox pilosa

Abstract: We analyzed the relationships between population size and reproductive characteristics in the perennial prairie forb Phlox pilosa , an obligate outcrossing butterfly-pollinated species. We examined 27 populations ranging in size from 9 to over 75,000 flowering ramets in two regions of the state of Iowa (eastcentral and northwest) in 1993 and 1994. We collected flowers from each population and scored them for pollen arrival to stigmas and number of pollen tubes. We collected fruiting ramets from each population at the end of the Phlox growing season and scored them for height, biomass, and reproductive variables, including the number of flowers initiated and opened and the number of capsules initiated and matured. In both years, population size was significantly correlated with the number of capsules matured per ramet. Differences between populations in capsule production were set primarily at the pollination stage. In 1993, pollen arrival to stigmas was significantly lower than in 1994 and was correlated with population size in eastcentral Iowa populations, indicating that lower reproduction in small populations that year was at least partially due to inadequate amounts of pollen being moved. In 1993, weather conditions likely depressed pollinator activity, but absolute capsule formation was high because of high flower production per ramet and high population densities. In 1994, when pollen arrival to stigmas was relatively high and unrelated to population size, outcross pollen movement was greater in larger populations. Increased efficacy of outcross pollen movement in 1994 may have resulted from lower flower production and less dense populations forcing greater pollinator movement between ramets or from variation between years in fine-scale spatial genetic substructuring of populations. Our results indicate that the viability of Phlox pilosa can be best ensured by protecting and creating populations of at least 1000–2000 flowering ramets.     

Effective Population Size in Red-Cockaded Woodpeckers: Population and Model Differences

Loss of genetic variability in isolated populations is an important issue for conservation biology. Most studies involve only a single population of a given species and a single method of estimating rate of loss. Here we present analyses for three different Red-cockaded Woodpecker ( Picoides borealis ) populations from different geographic regions. We compare two different models for estimating the expected rate of loss of genetic variability, and test their sensitivity to model parameters. We found that the simpler model (Reed et al. 1988) consistently estimated a greater rate of loss of genetic variability from a population than did the Emigh and Pollak (1979) model. The ratio of effective population size (which describes the expected rate of loss of genetic variability) to breeder population size varied widely among Red-cockaded Woodpecker populations due to geographic variation in demography. For this species, estimates of effective size were extremely sensitive to survival parameters, but not to the probability of breeding or reproductive success. Sensitivity was sufficient that error in estimating survival rates in the field could easily mask true population differences in effective size. Our results indicate that accurate and precise demographic data are prerequisites to determining effective population size for this species using genetic models, and that a single estimate of rate of loss of genetic variability is not valid across populations.